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Tuatara: Volume 26,Issue 2, November 1983

The Value of New Zealand Pollen and Spores as Indicators of Cenozoic Vegetation and Climates

page 37

The Value of New Zealand Pollen and Spores as Indicators of Cenozoic Vegetation and Climates

Keywords: ecology, New Zealand, palaeoclimatology, pollen, Quaternary, spores


Because of the widespread occurrence of organic sediments, New Zealand is ideally suited to the study of Cenozoic vegetation and climates by fossil spores and pollen. Primary descriptions and interpretations have already been made for most parts of the country and these are widely relied upon by a number of other disciplines.

In this paper we review the reasoning by which past vegetation and environments are reconstructed. Recent advances in plant ecology, historical ecology, eco-physiology, and palaeobotany, related to palaeoecology are reviewed.

The paper includes a comprehensive table giving the ecological ranges of plants found in the New Zealand pollen and spore record.


Quaternary pollen analysts have long had two complementary roles. The first is the reconstruction of past vegetation using pollen and spores preserved in sediment and unconsolidated sediments ranging from-calcareous loess to peat. The second is the ecologic interpretation of paleovegetation in terms of past environments, usually climate. The many pollen diagrams now available for New Zealand provide primary descriptions of past floras, vegetation and climates during the Tertiary and Quaternary as well as answers to more specific research questions e.g. identification of stadials/interstadials (Moar 1971, 1973; Moar & Suggate 1979; Mildenhall 1980).

Fossil pollen and spores are direct evidence only for past floras, with some indication of abundance (Faegri 1961). The reasoning by which the higher levels of ecological interpretation are reached are complex, difficult to define and usually subjective (cf. Rymer 1976). Difficulties of interpretation may or may not be able to be solved but this depends on the aim of the pollen analyst and the scale of the problem (Oldfield 1970).

page 38

The difficulties include:

  • 1) limits of specific pollen and spore identification, (often related to preservation).

  • 2) distinguishing between proximity and abundance of the source plants.

  • 3) differences in the area of vegetation represented by different pollen and spore types, defined as the pollen source area, and related to pollen productivity and dispersal power (Flenley 1973).

  • 4) changes over time in the pollen source area with change in floristics and structure of the local vegetation (Tauber 1965, 1967.)

  • 5) The conservative nature of many pollen types, especially those produced by rapidly speciating groups; this feature is an important problem in the New Zealand flora.

  • 6) The present-day ecology of plants represented in pollen assemblages is still often imperfectly known. The later part of this paper (Table 2) attemps to summarise current knowledge for New Zealand.

In this paper we review knowledge of present-day vegetation/pollen relationships to aid the critical appraisal of paleoecological data from New Zealand. The paper relies extensively on the studies of N.T. Moar and P. Wardle (Botany Division, DSIR) and D. T. Pocknall (N.Z. Geological Survey).

Representativity and Pollen Transport

Representativity is a measure expressing the relationship between species abundance and amount of pollen recorded at a particular location. Although the relationships can be expressed quantitatively for a particular site, (Davis 1963, Flenley 1973, Pocknall 1978, 1979), it is usually impossible to define the precise pollen source area. Moreover, differences in pollen productivity and dispersal power lead to different areas of vegetation being represented by the different pollen and spore texa at any time. The differences and variations between sites and over time have therefore led to representativity values being express qualitatively:

  • i Over-represented: pollen percentage is greater than the percentage of the source plants within the vegetation.

  • ii Well-represented: pollen percentage is approximately equal to percentage of the source plants within the vegetation.

  • iii Under-represented: pollen percentage is markedly smaller than percentage of the source plants within the vegetation, or zero.

These classes are based on the experience of the pollen analyst but are now usually supported by studies of the modern pollen rain in the same area as the fossil study. New Zealand examples include Moar (1969, 1970, 1971), Wardle (1971a), McKellar (1973), Myers (1973), Mildenhall (1976), Dobson (1976) and particularly Pocknall (1978, 1979, 1980).

Few texa indigenous to New Zealand have intrinsically high powers of pollen dispersion: Cyathea smithii-type, Dacrydium cupressinum, Nothofagus fusca type, Coprosma and Gramineae.

Other plants have lesser powers of pollen dispersal but nevertheless some 25 pollen and spore types are frequently encountered due to the widespread occurrence of the source plants across New Zealand:

Compositae, Phyllocladus, Podocarpus ferrugineus, P. spicatus, P. totara-type, Dacrydium bidwillii/biforme, Dacrydium colensoi, Nothofagus menziesii, Quintinia, Ascarina, Metrosideros, Weinmannia, Elaeocarpus, page 39 Griselinia, Pseudopanax, Myrsine, Pennantia, Leptospermum, Dicksonia, Cyathea dealbata-type, Phymatodes, Histiopteris, Monolete fern spores, Lycopodium spp. and Dacrycarpus dacrydioides. These types are usually the basis for interpreting and comparing pollen sequences even although they usually form only a small part of the total pollen flora preserved at any site. Consquently, there is often a broad similarity between fossil pollen diagrams from widely separated and floristically distinct areas in New Zealand.

The majority of New Zealand pollen and spore types are seldom found in modern assemblages unless the source plants are close to the site or water transport is involved. That is, these taxa are severely under-representative of the non-local vegetation but may be greatly over-represented if the plants grew at the pollen site. Most insect-pollinated trees, and the great majority of shrubs and herbs fall into this category. Wind-pollination is not necessarily a guarantee of good representation, e.g. Cyperaceae and Dacrycarpus.

A small group of plants exists for which pollen is virtually never found for instance the important hardwood trees Beilschmiedia tarairi and B. tawa (Macphail 1980). Such non-represented species are a “blind spot in New Zealand vegetation history, and pollen assemblages from Beilschmiedia- dominated forest are entirely misleading due to over-representation of local shrubs and distant strong pollen sources, usually conifers. Conversely severe under-representation can be an advantage in that even very low percentages of these taxa is good evidence for the local presence of the source, and for particular local environments if the source, plants have a well-defined ecologic range, such as Dactylanthus (Macphail and Mildenhall, 1980).

In addition to the intrinsic dispersal properties of individual pollen and spores types, the mode of pollen transport to the site also influences the magnitude of pollen deposition and size of the pollen source area. Studies in North-West Europe (Tauber 1965, 1967) suggest that in forest, pollen is transported to most sites in streams and surface run-off (Cw), through the spaces between tree trunks (Ct), above the canopy by wind (Cc) and by rain drops (Cr). Jacobson and Bradshaw (1981) have defined an additional pollen component (Cg) for pollen deposited vertically downwards from overhanging the sampling site. Tauber's model predicts that the relative proportions of these pollen components at a site is determined by the floristic and structural characteristics of the surrounding vegetation and nature of the site, particularly size and presence or not of inflowing streams. A corollary is that a change in site character, e.g. from a lake to a sedgeland, will alter the pollen source area and at such times, changes in vegetation seen via pollen analysis may have nothing to do with climate, (Ladd 1979).

Because spores and pollen are abundantly preserved in water-logged sediments such as peats and lake muds, basins where these sediments can accumulate continuously over long periods of time are favoured for pollen analytical studies. Fig. 1 shows the generalized relationship between the size of a site that has no inflowing stream and the relative proportions of pollen originating from different areas around the site: local- from plants growing within 20m of the basin edge; extra-local- from plants growing between ca. 20-300m from the basin edge; regional- growing at distances greater than ca. 300m (Jacobson and Bradshaw 1981 p. 82; Fig. 1). For shrub-covered peat bogs and basins of less than 1 ha, within forest, the major source of pollen will be local (Cg, Cw and Ct) Regionally derived page 40
Fig. 1 The generalized relationship between the size of a site that has no inflowing stream, the relative proportions of pollen originating from different areas around the site and level of vegetation reconstruction (after Jacobson and Bradshaw 1981)

Fig. 1 The generalized relationship between the size of a site that has no inflowing stream, the relative proportions of pollen originating from different areas around the site and level of vegetation reconstruction (after Jacobson and Bradshaw 1981)

pollen will penetrate into the sampling site (probably as Cr) but this long distance transported component will be dominated by locally derived types. The model predicts that as the dominated by locally dervied types. The model predicts that as the canopy opens up*, or the sampling site becomes incresingly distant from the dominant stratum, so extra-local and regional elements become increasingly well represented. Sub-tropical and tropical rainforest may however be an exception (Kershaw and Hyland 1975). Because pollen production of these rainforest trees is very low, domination of assemblages by local pollen does not occur and pollen deposition appears to be an accurate reflection of the regional vegetation.

All undisturbed hardwood, podocarp and beech forests and most forms of scrub in New Zealand are evergreen and closed (sensu Specht 1970). Most of the lakes utilized for paleovegetational research are small (less than 100m diameter) or if larger (Pocknall 1980) sampled close to the shore. Consequently, if the site is below the climatic treeline, the pollen sequences will preferentially record shifts in dominance with the local canopy and riparian communities. These shifts will be accompanied by some less well-defined indications of shifts in dominance elsewhere, particularly if the Cw component is important. So far there is no way to distinguish between a few strong pollen producers near the sampling site and, abundance of the same taxa in the regional vegetation. Consequently patterns in vegetation at the local and extra-local scale can only be detected by utilizing suites of sites dominated by local pollen.

The reverse situation occurs in low, open vegetation such as heath, page 41 grassland and herbfield. Local pollen influx may or may not be low but is usually augmented by pollen from the regional vegetation (Cr, Cc) and, at sites above the treeline, dominated by these taxa (Table 1). Here, occurrences of pollen from severely under-represented taxa that are restricted to cold climates, e.g. Donatia novae-zelandiae, Notothlapsi and Phyllachne may be the only definite indicators of the local enviroment whilst part of the long distance transported pollen is derived from the prevailing winds, much is derived from a preferential upslope transport of forest pollen types from lower elevations (Moar 1970, Heusser 1974, Macphail 1975; 1979, Hope 1976, Salgado-Labouriau 1979). With over-representation of distant, strong pollen dispersers, the boundaries between different vegetation types are blurred in the pollen record, particularly the precise location of the climatic treeline relative to the sampling site.

Modern pollen data supporting the above generalisatons are given in Table 1. These show the broad similarity between pollen assemblages from within Nothofagus temperate rainforest (Lake Hanlan, Shag Tarn) and those from sites in low, open vegetation (Denniston) or above the climatic treeline (Saddle Lakes). At Saddle Lakes Dacrydium bidwillii-type, Donatia novea zelandiae and Phyllachne are the only pollen evidence for the presence of the local alpine vegetation. The high percentage of Gramineae pollen at Denniston may reflect the local grassland but are more likely to be due to the occurrence of a few plants in the tank. The influence of close vegetation on pollen assemblages is more clearly seen in the moss samples from Lake Hanlan and Shag Tarn. At Lake Hanlan, the forest floor is virtually devoid of all but cryptogams and the moss sample is overwhelmingly dominated by Nothofagus fusca-type pollen. Despite the proximity of several emergent Dacrydium cupressinum trees, podocarp pollen is less that 5% implying very low wind velocities within the forest stand. At Shag Tarn, a moderately dense substratum of Coprosma spp. occurred and this is reflected in the relatively high shrub pollen percentages. Conversely the surface lake muds at both sites contained diverse palynofloras representing a range of local (riparian) and upland plant communities within the catchments.

Reconstruction of Paleovegetation

Knowledge of the relative pollen dispersal class of an individual pollen type, and identification of its probable mode of transfer to the sampling site, enable the pollen analyst to assess the relative abundance of that taxon in a fossil flora. Reconstruction of paleovegetation requires a second step, identification of the fossil plant communities.

A plant community can be defined as an assemblage of plants living together in an area (of any size) during the same time and in relatively stable proportions (Atkinson et al. 1968). Since fossil pollen sequences are an incomplete record of past floras and usually integrate the pollen production from a number of different communities, it is difficult directly to identify past plant communities. Instead one or more of three indirect approaches are adopted (Birks and Birks 1980):

1. Using statistical techniques to establish recurrent groups of fossil taxa, i.e. assemblages of fossil pollen and spores consistently occurring together, or numerically correlated in a series of samples within (and between) pollen sequences.

In New Zealand, Harris and Norris (1972) have used chi squared tests to identify significantly associated species pairs, using presence-absence data page 42
Table 1: Modern pollen spectra from Buller district, South Island
Percentages based on total woody species plus Gramineae. Percentages less than 1 % expressed as +
SiteLake HanlanShag TarnDennistonSaddle Lake LowerUpper
Location & Grid ReferenceKaramea Highway 3km south of Little Wanganui River Mouth S 17-18:547125.9km upstream of mouth Little Wanganui River S 17-18:61923225km north of Westport S24-264745Former glacial col at the head of the Little Wanganui River Valley. S 17-18:724164
Elevation (m)100 Nothofagus fusca closed forest with emergent Dacrydium cupressinum and subcanopy Weinmannia, Quintinia and Myrsine170 Nothofagus fusca closed forest with emergent Dacrydium cupressinum and subcanopy Weinmannia, Quintinia and Myrsine620 Grasses, subshrubs and adventives. 100m above edaphically depressed treeline, Nothofagus and Dacrydium spp.10801090 Mosaic of subalpine heath and sedgeland, including Dacrydium bidwillii and Olearia colensoi. 100m above local Nothofagus timber line.
Local Vegetation
surfacesurfacemoss in open tanksurfacesurface
Sample Typelake mudmosslake mudmossin abandoned townshiplake mudlake mud
Tall Podocarps5241327171722
Dryland Herbs212236+1012
Dacrydium bidwillii/biforme---+++33
Donatia, Phyllachne----+++
% Of Pollen Sum
page 43 on Quaternary pollen and spores preserved in deep peats from the Hauraki Plains. These were sorted into recurrent groups by an objective selection technique and assessed in terms of ecological significance. Other statistical methods, for instance Principal Components Analysis, have been successfully used eleswhere to determine current groups in modern and fossil assemblages, (Birks et al. 1975), Birks and Birks (1980).

2. Applying to past floras the ecological and phytosociological preferences of modern taxa.

If a species can be shown to occur in a limited number of associations, verifiable by objective ecological techniques, then the pollen of that species is assumed to indicate that association in the past. A related assumption is that the structure of the paleovegetation can be deduced by analogy when the past associations appear to agree with present-day well-defined associations.

3. Matching fossil pollen assemblages with modern assemblages from known vegetation types (the “finger-printing approach, Martin and Gray 1962, Wright 1967).

Approaches 2 and 3 depend on finding modern analogues so that the present can be extrapolated into the past. Both are common palynologic practice, at least implicitly, but are misleading if applied without caution to early Quaternary or pre-Quaternary assemblages. Their successful use requires sound knowledge of modern ecological preferences.

A serious problen in New Zealand is that numerous life-forms may be represented within the one pollen taxon (Rattenbury 1962), for instance.
Nothofagus fusca-type4 taxa
Metrosideros11 species
Coprosma45 species

Another problem is the difficulty of indentifying stable plant communities that are independent of climate, such as edaphically controlled Dacrycarpus semi-swamp forest and Westland “pakihi sedgelands of gley podsols (Rigg, 1962). Both communities include well-represented and palynologically distinct taxa.

Plant communities above the climatic treeline in New Zealand are usually well-defined floristically and uniform throughout the country. The tallest stratum is dominated by combinations of Compositae, Gramineae, Umbelliferae, Rubiaceae, Scrophulariaceae, Epacridacae and Liliaceate but structural and floristic analogues can also occur in exposed or semi-arid areas at lower elevations. Communities below the treeline are less easily defined. Although there are relatively few canopy species, mostly Podocarpaceae, Nothofagus spp., Weinmannia, Beilschmiedia, Elaeocarpus, Metrosideros and Quintinia. There is also a gradual replacement of mixed podocarp-hardwood stands by Nothofagus with increasing altitude, and latitude. Consequently these taxa can occur in any combination or proportion within the same climatic regime, (Holloway 1954, Robbins 1962, Nicholls 1976, Park and Walls 1978). Podocarp stands are widely recognized as having unstable population structures under present-day climates (Herbert 1980) and in some localities they are being competitively excluded by hardwoods and Nothofagus (Wardle 1963, Clayton-Green 1977).

page 44

Accordingly, Wardle (1964) has questioned the existence of stable forest associations and concluded that the component species have overlapping but largely independent distributions. These distributions are determined not only by differences in reproductive biology and habitat requirements (determining rates of migration and performance within the forest succession) but also by past events. The latter include the severe but geographically variable effects of Quaternary glaciation (Moar 1973, Burrows 1979) and volcanism (McKelvey 1963, McGlone and Topping 1977) as well as perennial phenomena such as landslipping, catastrophic weather and fires (Molloy 1969, Wardle 1978).

If canopy species have behaved individualistically through time, then it is clearly unwise to press analogies based on the modern vegetation too closely when inferring past associations. For example, Walker and Flenley (1979) have demonstrated that whilst some taxa in New Guinean ‘podocarp-hardwood-Nothofagus’ rainforest occur together with great fidelity through a variety of environmental vicissitudes, other taxa associated with them may behave independently when viewed on longer time scales during the late Quaternary. In New Zealand, this problem has to a large extent been avoided by interpreting past vegetation in terms of formations defined by pollen dominants and using minor, less well-represented taxa to suggest thermal or geographic variatons, (Moar 1971, 1973, McGlone and Topping 1973, 1977, Dobson 1978). This approach is appropriate to developing regional paleovegetational reconstructions but smaller scales of pattern have mostly yet to be interpreted.

Reconstruction of Past Environments

Reconstruction of past environments is usually based on both abiotic (lithology, geochemistry, site geomorphology, isotopes etc.) and biotic evidence, of which fossil pollen data is but one part. The most common method in pollen analysis is an extension of the ‘indicator species’ approach, the application backwards in time of known preference of (i) individual taxa with well-defined narrow environmental ranges or (ii) plant communities of taxa having similar ranges. Alternatively, statistical methods which relate pollen values to one or more environmental parameters using discriminant functions or regression equations may be used. A New Zealand example of discriminant function analysis relating groups of pollen and spores to temperature is provided by Harris et al. (1976). Transfer regression equations have been successfully used in a rainforest area of Chile by Heusser and Streeter (1980) and may be found applicable to New Zealand once the data for modern pollen spectra are adequate.

Much of the present information concerning the environmental tolerances of phytosociologically important species in New Zealand comes from compilations of available herbarium and field records: Ascarina lucida (McGlone and Moar 1977), Chionochloa rigida(Mark 1965), Dacrydium cupressinum (Franklin 1968), Leptospermum spp. (Burrell 1965), Metrosideros umbellata (Wardle 1971b), Nothofagus menziessii (Wardle 1967), N. solandri (Wardle 1970), Olearia colensoi (Wardle et al. 1970), Phyllocladus alpinus (Wardle 1969) and Weimannia racemosa (Wardle 1966). Agathis australis (Eckroyd 1982), Beilschmiedia tawa (Knowles and Beveridge 1982). Not all these taxa can be specifically identified by pollen. Ecological data on other equally important taxa is scattered and often unpublished.

page 45

Nevertheless, the information available does indicate that the majority of, if not all, phytosiciologically important trees have very wide environmental tolerances. In particular the taller podocarps are able to survive conditions which prevent regeneration by seed but, because of their longevity, are still able to influence the stability of rainforest communities over this period. Hardwood taxa appear to have shorter life-spans and would be excluded more rapidly under unfavourable conditions. However, the pollen of neither group provides a sensitive indication of climate except over long periods of time, suggested to be in excess of two thousand years.

Subcanopy trees and shrubs with more restricted distributions but with good representation are proving more rewarding as climatic indicators Ascarina lucida is one example (McGlone and Moar 1977) and trends in this taxon may serve to correlate early to middle Holocene pollen sequences across New Zealand. Dodonaea may be another (W. F. Harris pers. comm.). Because many of these taxa are under-represented, it is necessary to count large numbers of pollen or scan pollen slides for trace occurrences combined with a lower pollen count.

A remarkable example of the use of experimental plant physiology to interpret early postglacial sequences in the eastern South Island is given by Wardle and Campbell (1976) in their studies on frost-tolerance of plants found as pollen.

Table 2 lists pollen and spore types that (i) occur regularly and in some abundance in late Quaternary sediments or (ii) are rare but specifically identifiable or (iii) indicative of particular environments. Relative pollen dispersal classes are based on Dobson (1976), Pocknall (1978, 1979, 1980) and Macphail (unpub. results): Comments on the distribution and ecology take into account the available ecological literature (Wardle 1973, Orwin 1974) and extensive field observations by Wardle (1975, 1977) and D. R. McQueen but cannot take account of many anomalous distributions (Holloway 1954). Plant names and climate zones follow Allan (1961), Moore and Edgar 1970) and Wardle (1964).


There can be no doubt that pollen analysis has been spectacularly successful in providing primary descriptions of past vegetation and environments in New Zealand, particularly for the Late Quaternary period. Published descriptions are lacking only from North Auckland (M. S. McGlone pers. comm.) but all parts of the country will benefit from additional studies. At this scale trends in vegetation structure and climate parallel those in South-eastern Australia and Southern Chile (Macphail 1979, Heusser 1974).

Rather less is known of patterns in vegetation and climate on the smaller spatial and temporal scale. Where specific values for postglacial climates have been determined from fossil pollen assemblages, these are supported by macrofossils, e.g. Lintott and Burrows (1973), Soons and Burrows (1978), McGlone et al. 1978).

The problem of scale is similar to that confronting the ecologist studying modern plant communities. There is no reason to doubt that environmental factors determine pattern in vegetation on the geographic scale or over millenia, but it is difficult to assess the contribution of the same factors on individual stands or over periods of time less than several forest generations. Moreover, the ecological concepts underlying any extraction of environmental factors from fossil data are deterministic page 46
Table 2 (pp 46-57)
The present day distribution and ecology of plants represented by pollen and spores in New Zealand Quaternary and Recent deposits.
The use of the family or generic name only indicates that the pollen type includes more than one genus or species. Families, habit and distribution are from Allan (1961) and Moore and Edgar (1970). Ecology is from cited references and the authors' field observations. Climatic zones are in terms of Wardle (1964):
at 41 °Sat 45 °S
High Alpineover 1525mover 1220m
Lower Alpine1220-1525m915-1220m
Cool Temperate or Montane416-915m0-610m
Warm Temperate0-460mabsent
Pollen dispersal, from Pocknall (1979) and pers. comm.
LTD Long distance transportW Well to over-representedU Under-representedSU Severely under-representedNI No information
Pollen Taxon.FamilyHabitPollen DispersalEcology and Distribution
AcaciaMimosaceaeShrubs & treesLDTAustralia and introduced (Mildenhall 1972)
AcaenaRosaceaeHerbsUGrassland and open situations from coast to Subalpine.
Agathis australisAraucariaceaeLarge treesNIForest south to latitude 38°S, Warm Temperate to Montane, even at southern limit, and grows with Nothofagus menziesii(Ecroyd 1982).
Alectryon excelsumSapindaceaeSmall treeUWarm Temperate forest north of ca. latitude 42°S, usually on alluvium.
AmpereaEuphorbiaceaeShrubLDTAustralian (cf. Macphail 1979).
AristoteliaElaeocarpaceaeSmall tree: A. serrataUA. serrata: Temperate to Montane, scattered in intact forests but forming extensive areas of scrub after disturbance; frost tolerant.
Shrub: A. fruticosaA. fruticosa: Subalpine, drier to superhumid shrublands.
AvicenniaRhizophoraceaeTall shrubWEstuaries to latitude 38 °S, always in saline sites.
ArthropodiumLiliaceaeTall herbUOne sp. coastal rocks south to latitude 38 °S.
One sp. forest south of latitude 37 °S, Temperate.
Ascarina lucidaChloranthaceaeShrub-small treeWTemperate and Montane forests south of latitude 35 °S. Common in lowland forest in milder climate. Frost and drought intolerant.page 47
AsteliaLiliaceaeTerrestrial and epiphytic herbsUTemperate to Alpine in forest, scrub, grassland and herbfields throughout.
BeilschmiediaLauraceaeLarge treeSUOne sp. Warm Temperate forests south to 38 °S.
One sp. Warm Temperate forests south to 42 °S.
CalystegiaConvulvulaceaeHerbSUTemperate one sp. dunes, three spp. forest margins and open areas.
CarpodetusEscalloniaceaeShrubs & small treesUTemperate to Montane forests throughout; often on slips and in disturbed areas.
CasuarinaCasuarinaceaeShrubs & small treesLDTAustralia and introduced (Close et al. 1978 Macphail 1979)
Chenopodiaceae and Amaranthaceaeherbs and subshrubsW.LDTCoastal, especially saline habitats, throughout; extending inland in dry situations.
Collospermum hastatumLiliaceaeepiphytic & terrestrial tall herbsSULocally abundant in Warm Temperate forest south of ca. lat. 42 °S.
ColobanthusCaryophyllaceaeherbUSubalpine to Alpine, and rarely on compensating open sites to sea level.
Compositae (Liguliflorea)herbsNITemperate to Alpine, grassland and open areas.
Compositae (Tubuliflorae)herbs, shrubs & small treesUThroughout all plant communities at all elevations but less frequent in forest.
Abundant to dominant in coastal scrub, Subalpine scrub and Alpine herbfield.
CoprosmaRubiaceaeshrubs & small treesWCoastal to Lower Alpine; in forest, forest margins, scrub, grasslands. Tolerant of poorly drained as well as exposed habitats.
CordylineAgavaceaesmall trees, shrubsUTemperate on colluvium, river flats and base-rich swamps, 1 sp. in Montane and Subalpine forest in areas over 2000mm precipitation p.a.
CoriariaCoriariaceaeshrubWTemperate, Montane to Alpine throughout N.Z., often forming closed stands in successions colonizing recent soil.
Corynocarpus laevigatusCorynocarpaceaemedium treeSUCoastal Warm Temperate north of ca. lat. 44 °S; usually within or marginal to coastal forest (Stevenson 1978)
CruciferaeherbsUCommon on alluvium and distrubed ground, mainly coastal to open Alpine situations throughout, but 1 genus in forest.
CupressusCupressaceaemedium treeNIIntroduced and widely distributed as a windbreak tree.
Cyathea dealbataCyatheaceaetree fernUTemperate and Montane forests throughout.
C. medullarisCyatheaceaetree fernUTemperate forest, forest margins and disturbed forest; mainly North Island and west coast of South Island.page 48
C. smithii typeCyatheaceaetree fernWTemperate to Subalpine forest throughout; frost tolerant.
Cyathodes fasciculatusEpacridaceaeshrubUWarm Temperate-Montane; open situations in podocarp forest, common in Nothofagus forest, scrub and rocky places north of ca. lat. 44 °S.
CyperaceaeherbsWSwamps and wetlands at all elevations; throughout.
Dacrycarpus dacrydioidesPodocarpaceaetall treesUTemperate throughout, abundant or dominant in forests on wet alluvial soils and swamps. Extremely rare on Stewart Island (Wardle 1974), and not vigorous away from coast in Southland (Johnson 1972).
Dacrydium bidwillii/ biformePodocarpaceaelow shrubs & small treesUD. biforme: Montane & Subalpine south of lat.
36 °50'S in high precipitation forests and scrub.
D. bidwillii: Montane and Subalpine south of ca. lat. 36 °S; characteristic of gley podzol soils and bog, in scrub and openings in forest, descending to sea level along cold air drainage channels in south. Also present on well drained alluvium east of Southern Alps. Very frost tolerant (-20 °) (Wardle and Campbell, 1976).
D. colensoiPodocarpaceaemedium treeUMontane to Cool Temperate from ca. lat. 35 °S to 44 °S abundant on gley podzols and infertile swamps on coastal terraces in Westland. Always in high precipitation.
D. cuprsssinumPodoearnaceaetall treeW.LDTTemperate to Montane forest throughout; abundant over wide climatic range on all but very dry or waterlogged soils; intolerant of severe water stress, regenerates well after fire. Regeneration suggested to be cyclic, involving dicotylous trees as Weinmannia, Quintinia and Beilschmiedia (Poole 1937, Cameron 1954).
D. cf. intermediumPodocarpaceaesmall-medium treeWLowland forests in Northland, otherwise Montane to Subalpine throughout, particularly on infertile sites in west of South Island and Stewart Island.
Dactylanthus tayloriBalanophoraceaeroot parasiteUTemperate to Montane on wide range of hosts, mostly trees. Widely scatterd but locally common throughout North Island, probably present in Northwest Nelson (Macphail & Mildenhall 1980)
Dicksonia fibrosaDicksoniaceaetree fernUTemperate forests south of ca. lat. 37 °S.
D. lanataDicksoniaceaetree fernUTemperate to Montane from ca. lat. 35 °S to lat. 44 °S (on west coast of South Island) usually local, preferring warmer north-facing slopes.
D. squarrosaDicksoniaceaetree fernUTemperate to Montane throughout; abundant in lowland forests, particularly in shallow gullies and river flats, persisting after destruction of forest.page 49
Dodonaea viscosaSapindaceaeshrub-small treeUCoastal and, less commonly, inland from extreme north to ca. lat. 43°S; tolerant of exposed coastal situations, intolerant of frost.
Donatia novaezelandiaeDonatiaceaecushion plantSUAlpine on wet shallow soils; extended downslope onto gley podzol soils in cold air influenced sites.
DracophyllumEpacridaceaeshrubs & small treesSUCoastal to Alpine throughout; characteristic of Subalpine scrub and rocky Lower Alpine situations.
DrapetesThymelaceaesubshrubUUpper Subalpine and Alpine throughout.
Dysoxylum spectabileMeliaceaesmall to medium treeSULocally abundant in Warm Temperate forest from extreme north to ca. lat. 41 °30'S.
ElaeocarpusElaeocarpaceaemedium treesUTemperate and Montane forests throughout.
ElytrantheLoranthaceaehemiparasitic shrubsSUTemperate to Subalpine south of ca. lat. 36 °S; widely scattered and locally common on Quintinia, Pittosporum, Metrosideros and especially Nothofagus.
Epacridaceae (T-type)prostrate & erect shrubsSUCoastal to Subalpine throughout; in forest, scrub, grassland and bog.
EpilobiumOnagraceaeherbsSULowland to Lower Alpine throughout; usually in rocky or wet open situations.
EucalyptusMyrtaceaesmall-tall treeLDTAustralia and introduced (Close et al. 1978, Macphail 1979)
Eugenia maireMyrtaceaetreeNIWarm Temperate swamp forest south to lat. 41 °30'S.
EuphrasiaScrophulariaceaeherbsUTemperate to Alpine south of lat. 38 °.
Freycinetia banksiiPandanaceaelianeUCoastal and lowland from far north to ca. Marlborough in east and Fiordland in west of South Island; abundant in forest, and scrub especially near coast.
FuchsiaOnagraceaelianes, shrubs & small treesSUTemperate to lower Subalpine throughout; usually on forest margins or in openings.
GaultheriaEricaceaeshrubsUTemperate to Alpine south of lat. 37 °30', understorey in open forest; scrub and grassland. Important on volcanic scoria.
Geniostoma ligustrifoliumLoganiaceaetall shrubSUWidespread in coastal and Warm Temperate forests north of ca. lat. 41 °30'S.
GentianaGentianaceaeherbsSUTemperate to Alpine throughout, in grasslands, openings in heath and scrub, herbfield and fellfield.
GeraniumGeraniaceaeherbsSUTemperate to Lower Alpine grasslands throughout.page 50
GeumRosaceaeherbsSUMainly Subalpine and Lower Alpine grasslands but descending dowslope into cold air influenced open situations.
GleicheniaGleicheniaceaefernsUTemperate to alpine throughout; 2 spp. dominant on acid bogs.
Gramineae (Poaceae)sward-forming to tall tussockWTemperate to Alpine throughtout. Dominant in Alpine and semi-arid Temperate grassland communities, rare in undisturbed forest but locally extensive on lowland river flats, slips and sand landforms.
GriseliniaCornaceaeshrubs & samll trees frequently epiphyticUCommon in Temperate to Subalpine forests south of ca. lat. 35 °S; less commonly in subalpine scrub.
GunneraHaloragaceaeherbsSUTemperate to Subalpine throughout; common as a pioneer on stony river flats, river banks, slips, in grasslands and other open vegetation. Also in boggy situations within forest.
Gyrostemonaceaeshrub & small treesLDTAustralian (Macphail 1979)
HaloragisHaloragaceaeherbsUCoastal to Subalpine throughout; usually in open situations.
HebeScrophulariaceaeshrubSUCoastal to Alpine throughout; locally common and characteristic of seral scrub at low elevations, Subalpine scrub and grassland, Alpine herbfield and exposed rocky situations.
Hedycsrya arboresMornimiaceaesmall treeUTemperate forests from far north to Banks Peninsula on east, and along west coast of South Island; most common towards coast.
Histiopteris incisaPteridaceaefernNITemperate to Subalpine throughout; usually in shaded situations on forest margins, regenerates well after forest destruction in higher precipitiation areas.
HoheriaMalvaceaesmall treesSUTemperate to Subalpine, absent Stewart Island; usually marginal to forest and scrub but forming extensive scrub along river banks, on slips and avalanche pathways. Characteristic of wetter western districts but some species in lower precipitation areas.
H. glabrata, deciduous, resists -15 ° (Sakai et al. 1981).
HydrocotyleUmbelliferaeherbsUTemperate to Alpine throughout, usually in moister places.
IsoetesIsoetaceaeaquatic herbsNILocally common in higher elevation lakes and tarns, less common along upper reaches of rivers.
Ixerba brexioidesEscalloniaceaetreeNITemperate to Montane from lat. 35 °30'S to 38 °Spage 51
Knightia excelsaProteaceaetreeUTemperate south to Marlborough Sounds, often in late seral vegetaion.
Laurelia novaezelandiaeMonimiaceaetall treeSUWarm Temperate forests in North Island and north-west of South Island with scattered occurrences along west coast to ca. lat. 46 °S; characteristic of wet alluvial soils and base-rich swamps.
LeptospermumMyrtaceaeshrubs to medium treesUTemperate to Subalpine throughout; presistent in swamps and infertile soils but mainly in successional scrub following destruction of forest, on dunes, and in open low forest in drier areas.
LibertiaIridaceaetufted low to medium herbTemperate to Subalpine, forest floor and seral scrub.
LibocedrusCupressaceaesmall to medium treesU1 sp. Lowland forest from ca. lat. 35 °S to N.W. Nelson; 1 sp: upper Montane and Subalpine forests south of ca. lat. 36 °S, frost tolerant to -13 ° (Sakai et al. 1981). Pure stands of catastrophic origin (Clayton-Greene 1977).
Liparophyllum guniiGentianaceaeherbsSUMontane to Alpine south of ca. lat. 39 °S; local in herbfields and bogs. Near sea level in far south.
Lophomyrtus -typeMyrtaceaeshrubs & small treesUTemperate to Subalpine throughout; forest understorey and margins.
Loranthus micranthusLoranthaceaehemiparasitic shrubSUTemperate throughout; common on a wide range of hosts including Carmichaelia, Coprosma, Hoheria, Leptospermum, Cyathodes, Lophomyrtus, and Podocarpus.
Lycopodium australianumLycopodiaceaeherbUMontane to Lower Alpine, south of ca. lat. 38 °30'S; usually in stony grassland, low heath, and bog. Descends to lower elevations in exposed or cold air influenced situations.
L. deuterodensumLycopodiaceaeherbUWarm Temperate from far north to ca. lat. 38 °S. Isolated occurrences in Marlborough.
L. fastigiatum-typeLycopodiaceaeherbWMontane to Lower Alpine throughout; usually in grassland and bog descending into lowlands in extreme south.
L. scariosumLycopodiaceaeherbWTemperate to Montane south of ca. lat. 38 °S; local in open forest situations.
L. varium-typeLycopodiaceaeherbs, sometimes epiphyticWCommon in Temperate and Montane forests throughout.
Lygodium articulatumSchizaeceaelianeNIWarm Temperate south to Lat. 38 °S.
MelicytusViolaceaesmall treeUTemperate to Subalpine throughout, forest openings and seral forest.page 52
MetrosiderosMyrtaceaelianes, shrubs, small to tall trees: epiphytic juvenilesWTemperate to Subalpine throughout; taxon including important canopy species of lowland forest. Involved in cyclical regeneration with Dacrydium cuypressinum(Wardle 1971).
Metrosideros excelsaMyrtaceaetreeWWarm Temperate, coastal south to lat. 38 °S, generally on bluffs and sea cliffs.
M. robustaMyrtaceaetall treesWTemperate to Montane from extreme north to ca. lat. 42 °30'S on west coast of South Island; a canopy tree in lowland forest.
Microseris-typeCompositaeherbsWNative and introduced; characteristic of grasslands and disturbed ground.
Mida salicifoliaSantalaceaeshrub-small treeSUTemperate to Montane forest from extreme north to north-west coast of South Island. Sparsely distributed except in north of its range.
Monolete Fern Sporesnumerous families of Order FilicalesfernsW,U,SUTemperate to Alpine throughout; characteristic of the understorey in lowland and montane forests, colonizing slips, river banks and other damp shady habitats.
MuehlenbeckiaPolygonaceaelianes, shrubsUTemperate to Subalpine throughout; characteristic of exposed coast, disturbed forest; forest margins, river flats and grasslands.
MyriophllyumHaloragaceaeaquatic herbsWTemperate to Subalpine throughout; tolerant of saline to freshwater, stagnant to flowing conditions.
MyrsineMyrsinaceaeprostrate shrubs to treesUTemperate to Alpine throughout; common in forest, forest margins, scrub on poorly drained sites and extending into Alpine grasslands and rocky areas.
NestegisOleaceaetreesNITemperate, North Auckland to 41 °30'S.
Nothofagus fusca-typeFagaceaetreeW,LDTTemperate to Subalpine throughout, often forming treeline; dominant or potential dominant of temperate Montane and Subalpine forest in precipitation from 1000mm to over 7000mm p.a. Fertility requirements vary with the individual taxa. N. fusca > (N. menziesii) > N. truncata > N. solandri var. cliffortioides (Adams 1976); the latter frost tolerant to -13 ° (Sakai et al 1981).
N. menziesiiFagaceaetreesUMontane and Subalpine south of lat. 37 °S on North Island and along west coast of South Island, often forming the treeline. Dominant of Cool Temperate forests in south of South Island.
NotothlaspiCruciferaeherbsSUAlpine-subalpine screes and rocky areas south of ca. lat. 41 °S.page 53
OphioglossumOphioglossaceaeherbsUOpen grassy areas throughout.
Pennantia corymbosalcacinaceaesmall treeWTemperate forests south of ca. lat. 35 °S; abundant on alluvial and colluvial soils.
PhormiumAgavaceaetall tussockUTemperate to Subalpine throughout; abundant in coastal scrub and fertile swamps, river and lake margins and in subalpine forest, scrub and grassland.
PhyllachneStylidiaceaecushion plantsSUSubalpine to Alpine herbfields, fellfield and rocky areas south of ca. lat. 39 °S.
PhyllocladusPodocarpaceaeshrubs & small treesW,LDT2 spp. scattered in Temperate forest of North Island and north-west coast of South Island; P. alpinus characteristic of Subalpine forest and scrub, reaching sea level in south on infertile, gley podzol soils and bogs. Frost tolerant to -18 ° (Sakai et al 1981). Locally abundant after fires.
PhymatodesPolypodiaceaeferns, often epiphyticWTemperate to Montane throughout; locally abundant as an epiphyte in forest, occasional amongst rocks on coast and at high elevation.
PinusPinaceaetreeW,LDTIntroduced throughout; widely planted for forestry in North Island and naturalized in open areas up to Montane zone.
PimeleaThymeleaceaeshrubsSUTemperate to alpine throughout; characteristic of semi-arid, Subalpine and Alpine grasslands.
PittosporumPittosporaceaelianes, shrubs & small treesWUTemperate to subalpine throughout; often seral.
PlagianthusMalvaceaeshrubs to small treesSUTemperate to Montane throughout; characteristic of salt marshes, estuaries, river flats, and forest on alluvial flats.
PlantagoPlantaginaceaeherbsUTemperate to Alpine throughout; rare within forest and scrub but characteristic of alpine herbfield and grasslands; usually in damp situations.
P. lanceolataPlantaginaceaeherbW,LDTIntroduced throughout; widespread as pastoral weed.
Podocarpus ferrugineusPodocarpaceaetall treesW,UCommon to locally abundant in Temperate and Montane forests throughout.
Frost tolerant to -10 ° (Sakai et al 1981).
P. spicatusPodocarpaceaetall treesW,U,Common in Temperate forest on fertile soils throughout, but rare on Stewart Island. Prefers warmer, drier situations.
P. totara-typePodocarpaceaeprostrate shrubs to tall treesW,U4 spp. Temperate to Subalpine, common lowland river flats, in forest and high elevation scrub.page 54
PolygonumPolygonaceaeherbsSUNative (north of ca. lat. 44 °S) and introduced (throughout); common in lowland swamps, pastures and along river banks.
PomaderrisRhamnaceaeshrubsULocal on dry hills in North Island; most species confined to Northland.
P. cf. apetalaRhamnaceaeshrub to small treeU,LDTIsolated in coastal situations on North Island; Australia (Macphail 1975)
PortulacaceaeherbsNITempoerate to Alpine, generally subaquatic but also on screes.
PotamogetonPotamogetonaceaeaquatic herbUCoastal and lowland; common in brackish and freshwater ponds, swamps and bogs. Rare at higher elevations.
PseudopanaxAraliaceaeshrubs & small treesUTemperate to Subalpine throughout; common in forest, forest margins and scrub.
PseudowinteraWinteraceaeshrubs & small treesSUTemperate to Lower Subalpine throughout; in forest, forest margins and scrub of higher precipitation; abundant on recent soils and often forming an extensive scrub after forest destruction.
Pteridium esculentumPteridaceaefernUTemperate to Montane throughout; characteristic of disturbed areas, forest margins at lower elevations and dry rocky places at high elevation. Rare within undisturbed vegetation.
PterisPteridaceaefernNIWarm Temperate south to ca. lat. 42 °S, forest openings and stream banks.
QuintiniaEscalloniaceaesmall treeWLocal in Montane forests of North Island but abundant in Temperate and Montane forest on the west coast of South Island; important subcanopy species with podocarps in central Westland “beech gap.
Ranunculaceaeherbs, lianesSUTemperate to Alpine throughout; native and introduced. Characteristic of disturbed, stony places in lowlands and coast, otherwise at higher elevations in grassland and herbfield. Includes Temperate forest lianes.
RestionaceaesedgeUTemperate to Subalpine; in salt marshes, and subalpine bogs throughout.
Rhopalostylis sapidaPalmaepalmSUWarm Temperate, coastal and adjacent lowlands on North Island, southward to Banks Peninsula and Greymouth in South Island. Abundant only near coast, often persisting after destruction of forest.
Ripogonum scandensSmilacaeaelianeUAbundant in Temperate podocarp-dicotylous forest.
RubusRosaceaelianes, scrambling shrubWTemperate and Montane throghout; abundant in disturbed forest, forest margins and along river flats. 1 sp. in semi-arid grassland.page 55
RumexPolygonaceaeherbWNative and introduced, lowland to montane throughout; characteristic of beaches, pastures, disturbed ground and grasslands.
RuppiaRuppiaceaeherbUBrackish to fresh water, Temperate - Montane.
Salicornia australisChenopodiaceaeherbWTemperate, coastal only, saline herbfields south to Foveaux Strait.
Schefflera digitataAraliaceaesmall treeUTemperate throughout; important species in moist gully forest.
SchizeleimaUmbelliferaeherbNIGenerally Alpine but occasionally Subalpine to Temperate.
Sparganium subglobosumSparganiaceaegraminoidWTemperate aquatic.
StellariaCaryophyllaceaeherbsSUTemperate to Alpine; grassland and rocky areas.
Tetrapathaea tetrandraPassifloraceaelianeUTemperate to lat. 44 °S, generally in open forest, or at forest margin.
Toronia toruProteaceaeshrubNIWarm temperate from lat. 30 °S to lat. 38 °S. Open forest and scrub.
Trilete Fern Sporesvarious families of order FilicalesfernsSUThroughout, in all but driest plant communities.
Tupeia antarcticaLoranthaceaehemiparasiteNITemperate to montane throughout.
Typha orientalisTyphaceaetall herbWTemperate swamps throughout.
Umbelliferae (excluding Hydrocotyle Schizeleima)herbsSUTemperate to Alpine throughout; forest margins, grasslands and open communities.
UrticaUrticaceaesuffruticose herbNITemperate, and subantarctic on richer colluvium.
WahlenbergiaCampanulaceaeherbsSUTemperate to Alpine throughout; characteristic of Alpine herbfield, and grassland.
WeinmanniaCunoniaceaemedium treeUTemperate to Subalpine south of ca. lat. 37 °S. Very abundant canopy tree in lower forests on west coast of South Island. Occurs on a wide range of soils; involved in cyclic regeneration with Quintinia and Dacrydium cupressinum (Wardle 1966), and forms part of successions on new ground. Intolerant of drought and heavy frost. Regenerates well after fire.
page 56 whereas succession in the shorter-term may be probabilistic, resulting in any one of a large number of possible species combinations that are ecologically compatible (Webb et al. 1972). Such alternative sucessions have been demonstrated for Tasmanian rainforest (Jackson 1968, Bowman and Jackson 1981) and is suspected for tropical rainforest in Queensland (Webb et al. ibid).

Whether New Zealand rainforest comprises independently distributed species as suggested by Wardle (1964) or whether stable associations and unifying ecological trends exist, it is clear that even a single cycle of regeneration or establishment is too long for any individual to observe directly. Processes in forest succession may be inferred by integrating observations made on suites of stands, as Wardle (1980) has done using moraines in the Westland National park. Alternatively, well-dated long pollen sequences can provide detailed evidence for successional trends in the one place, (Macphail, in prep.) Additional insights are being provided by historical records (Beever 1981). Macrofossil assemblages (e.g. Molloy and Cox 1972, Campbell et al. 1973, Drake and Burrows 1980) as well as plant population studies (e.g. Clayton-Green 1977, Wardle 1978) and tree nutritional data (Adams 1976), have also made significant contributions to our knowledge of successions past and present.

Many fundamental questions remain unaddressed. What is certain is that New Zealand's abundant, deep and varied organic deposits will enable the specific paleoecological queries to be isolated and answered, be they botanic, edaphic or climatic.


We are indebted to many for their help in the production of this paper, which arises from work during a Post-Doctoral Fellowship help by M. K. Macphail at the Botany Department, Victoria University of Wellington. Terese Hughes (University of Tasmania) and Maureen Cooper (Victoria University of Wellington) for their considerate and considerable typing efforts. D. C. Mildenhall and D. T. Pocknall (N.Z. Geological Survey), and N. T. Moar and M. S. McGlone (Botany Division, DSIR) aided by critically assessing the manuscript.


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1 Present Address: ESSO, Australia Ltd, G.P.O. Box 4047, Sydney 2001, Australia.

* Deciduous forests are a special case since sites within a forest stand become fully exposed to the regional pollen rain (if any) during winter. Greater wind velocites within the trunk space lead to an enhanced dispersal of pollen from those subcanopy species which flower before the forest trees come into leaf, e.g. Corylus.